Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 1 of 15

First published on the Flora Mesoamericana Website, 29 Dec. 2011.

61. DILLENIACEAE Family description and key to genera by C.A. Todzia y G. Aymard C.

Lianas or thick woody vines more than 40 cm in circumference., climbers, or shrubs or small trees with tortuous branches (). Leaves alternate, rarely opposite, simple, entire or dentate, usually with numerous parallel lateral nerves, frequently coriaceous and scabrous, glabrous or pubescent with sclerified or silicified hairs, simple or fasciculate; stipules absent, or wing-like and adnate to the petiole, deciduous. Inflorescences axillary or ramiflorous, cymose, racemose or the flowers solitary. Flowers bisexual or polygamous (and the androdioecious), white or yellow, hypogynous; sepals (3-7) frequently 5, spirally imbricate, persistent; petals 2-6, imbricate, often crumpled in bud, deciduous, usually thin and delicate; stamens usually numerous, free, persistent, the anthers opening by longitudial slits or apical pores; carpels 1-20, free or connate along the ventral side; style usually slender and elongate, with terminal, often capitate stigma; ovules 1-many in each carpel, apotropous. Fruit a follicle, capsule or berry, dry and dehiscent, or, as in , indehiscent and then included in a more or less fleshy calyx; seeds arillate; endosperm copious, carnose, the embryo very small. 10 gen., c. 350 spp., mostly in tropical and subtropical regions, especially Australasia. Bibliography: Kubitzki, K. Mitt. Bot. Staatsamml. München 9: 1-105 (1971).

1. Plants cultivated; trees; flowers 7-20 cm broad. 3. Dillenia 1. Plants native; usually or scandent shrubs, rarely trees (Curatella); flowers 0.3- 1.5 cm broad. 2. Leaves scabrous; carpels pyriform, 1-5, the ovules several per carpel; seeds 1-few, the aril lacerate. 6. 2. Leaves scabrous to smooth; carpels glabose, 1-2, the ovules 2 per carpel; seeds 1-2, the arile essentially entire. Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 2 of 15

3. Sepals unequal, the inner 2 larger and enclosing the fruit; fruit 1-seeded. 2. Davilla 3. Sepals nearly equal, not enclosing fruit; fruit with 1-2 seeds per carpel. 4. Small trees or shrubs; leaves stellate-pubescent on both sides, scabrous. 1. Curatella 4. Lianas; leaves glabrous or pubescent with simple hairs, smooth. 5. Leaves acute to acuminate at apex, lanceolate, elliptic, obovate, dentate; style 1; fruit globose. 4. 5. Leaves rounded at apex, suborbicular-ovate, obovate; entire or coarsely undulate; styles 2; fruit didymous. 5.

1. Curatella Loefl.. Por C.A. Todzia.

Small trees or shrubs; trunk short and branches tortuous, with harsh, rough pubescence, Leaves alternate, coriaceous, scabrous, pubescent with stellate hairs, incrusted with silica, pinnately veined. Inflorescences axillary, paniculate, many-flowered; flowers bisexual, actinomorphic, small, white; sepals 4-5, spreading, persistent; petals 4-5, caducous; stamens many; anthers oblong, longitudianlly dehiscent; carpels usually 2, globose, fused basally; ovules 2, erect, basal; styles filiform; stigmas peltate. Fruits capuslar, usually bilobed, globose, connate to base; seeds (1-)2 per carpel, smooth, almost completely surrounded by a membranaceous aril. 1 sp. American tropics.

1. Curatella americana L., Syst. Nat., ed. 10, 2: 1079 (1759). Neotype (designated by Todzia y Barrie, 1991): Venezuela, Williams y Alston 97 (LL!). Illustr.: Gallardo-Hernández, Fl. Veracruz 134: 7, t. 1 (2001). Trees or shrubs 2-10 m. Stems torulosus, with grayish to reddish flaking epidermis. Leaves 8-20(-35) × 5-10(-15) cm, short-petiolate, ovate to elliptic, rounded at apex, decurrent at base, craspedodromous with 10-22 pairs of lateral veins, scabrous and sparsely pilose above, sparsely to densely pilose beneath with stellate hairs, margin uneven and slightly undulate. Inflorescences axillary racemes or panicles, 4-9 cm. Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 3 of 15

Flowers 7-12 mm broad, pedicellate; sepals 5-7 mm, obovate, pubescent; petals white to pink, 5-7 mm, obovate; stamens 80-100. Capsules 4-6 mm, coriaceous, pubescent; seeds c. 4 mm, black, nitid, a white aril, nearly enveloping the seed at maturity. Common in secondary forests and savannas. T (Ventura 20628, MO); Ch (Breedlove 26678, MO); C (Chan 4702, MO); B (Davidse y Brant 32744, MO); G (Contreras 9786, MO); H (Williams y Molina R. 10666, MO); ES (Davidse et al. 37132, MO); N (Stevens 7612, MO); CR (Liesner y Lockwood 2434A, MO); P (Nee 10787, MO). 0-900 m. (C. Mexico, Mesoamerica, Colombia, Venezuela, Guayanas, Perú, Bolivia, Brazil, Paraguay, West Indies.) Kubitzki (1971) states that no material of Curatella could be found in the Linnaean herbarium. 2. Davilla Vand. Por C.A. Todzia.

Small climbing shrubs or woody lianas. Leaves petiolate, winged to narrowly winged, frequently scabrous, often pubescent with simple hairs only. Inflorescences terminal panicles or axillary in the uppermost leaves nodes. Flowers bisexual, actinomorphic, yellow; sepals 5, unequal, the 2 inner ones larger, very concave, accrescent, becoming coriaceous, enclosing the fruit and simulating a globose capsule; petals 3-6, obovate, deciduous; stamens numerous; carpels 1-2, free, 1-loculate; ovules 2, erect, basal; styles clavate; stigmas peltate, emarginate. Fruits globose, indehiscent or rupturing irregularly; seed 1, smooth, with the aril completely enveloping the seed at maturity. Approx. 20 spp. Mexico to Brazil.

1. Larger sepals pubescent, scabrous; leaves scabrous above softly adpressed pilose beneath. 1. D. kunthii 1. Larger sepals glabrous or laxly pilose, smooth; leaves smooth to scabrous above, pubescent beneath only along the veins or glabrous. 2. D. nitida

Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 4 of 15

1. Davilla kunthii A. St.-Hil., Pl. Usuel. Bras. sub t. 22 (1824 [1825]). Holotype: Venezuela, Humboldt & Bonpland 252 (P). Illustr.: Gallardo-Hernández, Fl. Veracruz 134: 11, t. 2 (2004). Davilla aspera (Aubl.) Benoist, Tigarea aspera Aubl. Lianas or small trees, with exfoliating bark; young branches sparsely strigose. Leaves 6-18 × 3-8 cm, elliptic to suborbicular, obtuse to short-acuminate at apex, rounded to cuneate at base, with 10-29 prominent lateral nerves, scabrous above, softly villous beneath. coriaceous, margins dentate, but often revolute; petioles 1-3 cm. Inflorescences terminal or axillary; panicles 6-15 cm, ferrugineous-villous. Flowers 8-15 mm broad; inner sepals broadly ovate to circular, densely hirsute, orange; petals 6-9 mm, obovate, yellow; stamens 50-85; carpel 1, glabrous; seed 3-4 mm. Premontane wet forests, savannas, disturbed moist forests, gallery forests. T (Cowan 1998, MO); Ch (Breedlove y Almeda 57804, MO); C (Martínez S. 11265, MO); QR (Cabrera y Cabrera 4176, MO); B (Davidse y Brant 32455, MO); G (Contreras 6848, MO); H (Saunders 152, MO); N (Stevens 7914, MO); CR (Davidse y Herrera 26299, MO); P (Nee 9134, MO). 0-1200 m. (C. Mexico, Mesoamerica, Colombia, Venezuela, Guayanas, Ecuador, Perú, Bolivia, Brazil, Trinidad.)

2. Davilla nitida (Vahl) Kubitzki, Mitt. Bot. Staatssamml. München 9: 95 (1971). Tetracera nitida Vahl, Symb. Bot. 3: 70 (1794). Holotype: French Guiana, Rohr s.n. (C). Illustr.: Standley y Williams, Fieldiana, Bot. 24(7): 5, t. 1 (1961). Davilla aspera (Aubl.) Benoist var matudae (Lundell) L.O. Williams, D. matudae Lundell, D. multiflora (DC.) A. St.-Hil., Tetracera multiflora DC. Lianas with exfoliating bark; young branches glabrous. Leaves 5-18 × 3-8(-9) cm, elliptic, oblong to lanceolate, rounded, acute, or apiculate at apex, attenuate to rounded, sometimes oblique, at base, with 6-13(-17) lateral nerves, smooth to slightly scabrous above, pilose beneath only along veins, coriaceous, margins entire to dentate; petioles 1-3 cm. Inflorescences 7-20 cm, terminal or axillary, sparsely pubescent. Flowers 7-12 mm broad; inner sepals broadly ovate, glabrous, orange; petals 4-6 mm, obovate; stamens 40- 50; carpel 1, glabrous; seeds 2.5-4 mm, with a white aril. Disturbed forest and roadsides, gallery forests. Ch (Breedlove y Smith 21834, MO); B (Gentry 8236, MO); G (Contreras Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 5 of 15

9345A, MO); N (Stevens 19974, MO); CR (Hammel 11454, MO); P (McPherson 6951, MO). 0-800 m. (S. Mexico, Mesoamerica, Colombia, Venezuela, Guayanas, Ecuador, Perú, Bolivia, Brazil, Cuba, Jamaica.)

3. Dillenia L. Por. C.A. Todzia.

Trees or shrubs. Leaves spirally arranged, pinnately nerved, entire or dentate; petioles narrowly to broadly alate. Inflorescences racemose, or flowers solitary, either axillary or terminal. Flowers bisexual, pedicellate; sepals usually 5, free, imbricate, persisten; petals usually 5, caducous; stamens many, either all of about the same length or in 2 more or less sharply separated groups of different lengths; anthers linear, opening by longitudinal slits or apical pores; carpels 4-20, arranged in a whorl around a broadly narrowly conical protruding part of the receptacle, mostly free; ovules 4-80 per carpel. Fruits consisting of gynoecium and calyx, dehiscent; carpels spreading; seeds arillate. C. 60 spp. Madagascar to the Fiji Islands, Australia. Literature: Hoogland, R.D. Blumea 7: 1-145 (1952).

1. Leaves with 30-40 pairs of lateral veins; flowers solitary, 10-20 cm broad; petals 7-9 cm; fruits 8-10 cm in diameter. 1. D. indica 1. Leaves with 8-17 pairs of lateral veins; flowers in few-flowered racemes, 7-9 cm broad; petals 3-5 cm; fruits 2-2.5 cm in diameter. 2. D. suffruticosa

1. L., Sp. Pl. 535 (1753). Lectotype (designated by Hoogland, 1952): Rheede, Hort. Malab. 3, 39, t. 38-39 (1682). Illustr.: not found. Evergreen trees to 30 m; trunk shallowly grooved and with many knots; young branches villous. Leaves 10-35 × 5-15 cm, oblong, with 30-40 pairs of lateral veins, acute to acuminate at apex, rounded to attenuate at base, the margin dentate with veins ending in apex of teeth, smooth, strigose along veins beneath; petioles 2.5-10 cm, glabrous to sparsely strigose. Flowers 10-20 cm broad, solitary, terminal; pedicels 4-8 cm; sepals 5, in open flower forming a cup-shaped calyx, yellowish-green; styles ca. 2.5 cm. Fruits Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 6 of 15

indehiscent, globose, 8-10 cm in diameter, yellowish-green. Cultivated in wet to seasonally dry areas. H (Sierra 69, MO); N (Araquistain 3498, MO); CR (Stevens 13591, MO); P (Croat 5920, MO). 0-1000 m. (Native to Sumatra, Java, Borneo, India to China; widely cultivated.)

2. (Griff.) Martelli, Malesia 3: 163 (1886-1890 [1887]). Wormia suffruticosa Griff., Not. Pl. Asiat. 4: 706 (1854). Syntype: Malaysia, Griffith s.n. (K). Illustr.: not found. Trees or shrubs 2.5-10 m; young branches sparsely pubescent. Leaves 12-30(-50) × 5.5-18 cnm obovate to broadly elliptic, rounded to acute at apex, decurrent along petiole, with 8-17 pairs of lateral veins extending to margin, smooth, glabrous above, strigose along midvein and axils of lateral veins, coarsely dentate; petioles 2-3 cm, sparsely strigose, broadly alate. Inflorescences 10-20 cm, lax racemes, with 4-8 flowers. Flowers 7-9 cm borad; sepals 1-1.5 cm, elliptic to ovate, glabrous; petals 3-5 cm, yellow, thin, glabrous; stamens c. 60; carpels 6-8, glabrous, each with 1-4 ovules; styles c. 1 cm. Fruits 2-2.5 cm, dehiscent, opening nearly flat, red; seeds obovoid, enclosed by a scarlet membranous aril. Cultivated. H (Meyer 9946, MO); ES (Berendsohn y Berendsohn WB 0030, MO); CR (Solomon 19146, MO); P (Schmalzel 270, MO). (Native to Indo- Malaysia; widely cultivated in the tropics.)

4. Doliocarpus Rol. Por C.A. Todzia y G. Aymard C.

Woody lianas or scandent or sarmentous shrubs. Leaves smooth, pubescnet with simple hairs only, dentate to subentire; petioles narrowly alate; stipules absent. Inflorescences axillary, ramiflorous, many-flowered, paniculate or fasciculate or glomerate. Flowers bisexual, actinomorphic; sepals (3-)5-6, unequal, persistent; petals 3-5, obovate, caducous; stamens numerous, the filaments filiform, the anthers small, longitudinally indehscent; carpels 1, 1-locular; ovules 2, anatropous; style terminal, stigma peltate. Fruit baccate, dehiscent or indehiscent, globose; seeds 1-2, arillate, the embryo small. Approx. 26 spp. Tropical America, chiefly Brazil. Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 7 of 15

1. Flowers and fruits sessile, subsessile, or less that 4-6 mm; plants uncommon. 2. Leaves c. 23 cm, pubescent beneath, tertiary veins raised. 1. D. dressleri 2. Leaves 6-15 cm, glabrous, tertiary veins not raised. 2. D. brevipedicellatus 1. Flowers and fruits on peduncles 3-30 mm; plants common. 3. Peduncles 2-8-flowered. 5. D. multiflorus 3. Peduncles generally 1-flowered. 4. Carpel glabrous, style c. 1 mm; fruit glabrous. 3. D. dentatus 4. Carpel pilose, styel 4-8 mm; fruit strigose. 5. Leaves punctate; sepals pubescent externally; fruits 1-1.3 cm in diameter, with strigose hairs less than 0.5 mm. 4. D. major 5. Leaves not punctate; sepals glabrous externally; fruits 1.5-2 cm in diameter, with strigose hairs 1-2 mm. 6. D. olivaceus

1. Doliocarpus dressleri Aymard, Novon 3: 317 (1993). Holotype: Panama, Dressler 3468 (MO). Illustr.: not found. Lianas leñosas; ramas grisáceas, glabras. Hojas 22 × 13 cm, cartáceas a subcoriáceas, oblongo-obovadas, glabros por la haz, pilosos por el envés, principalmente en el nervio principal y nervios secundarios, venación terciaria reticulada, discretamente areolada; base redondeada; márgenes subcrenados; ápice obtuso; pecíolos 2.8 cm, canaliculados, muy robusto en la base, balsalmente adpreso-pubescente. Inflorescencias glomeruladas; pedúnculos 4-5 mm; no ramificados, densamente adpreso-pubescentes; Flores con los sépalos 5, 5-8 × 4-6 mm, obovado-elípticos, seríceos externamente, laxamente seríceos internamente, principalmente en la parte media; pétalos no vistos; estabres 50-60, los filamentos glabros o pubérulados, las anteras c. 0.5 mm; ovario densamente híspido. Frutos 9 mm de diámetro, globosos, adpreso-pubescentes. Bosques muy húmedos. P (Dressler 3468, MO). 700-800 m. (Endémica.)

2. Doliocarpus brevipedicellatus Garcke, Linnaea 22: 47 (1849). Holotype: Surinam, Kegel 1186 (GOET). Illustr.: not found. Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 8 of 15

Lianas; young branches glabrous. Leaves 6-15 × 2.5-7 cm, lanceolate to elliptic, acuminate at apex, attenuat at base, with 6-9 pairs of lateral veins, the teritary veins not raised, smooth, strigose beneath along veins, coarsely dentate along c. upper 1/2 of margin; petioles 0.5-1 cm, strigose, very narrowly alate. Inflorescences 0.5-1 cm, fascicles of densely compacted 2-4-flowered racemes or flowers solitary; peduncles 1-3 mm, sparsely pilose. Flowers 3-5 mm broad; sepals unequal, outer ones 1.5-2 mm, inner ones 3-3.5 mm, ovate-elliptic, sparsely pilose outside, glabrous within; petals 2-3 mm, white; stamens 30-40; carpels 1, glabrous, style c. 1 mm. Fruits 5-6 mm in diameter, glabrous, red, dehiscent; seeds 1, with a white aril. Lowland wet forests. 0-400 m. N (Sandino 4886, MO); CR (Quesada 422, MO); P (Steyermark y Allen 16756A, MO). (Mesoamerica, Venezuela, Surinam, Bolivia, Brazil, West Indies.) Only one subspecies, subsp. brevipedicellatus, occurs in Mesoamerica; subsp. hilarianus is restricted to Brazil.

3. Doliocarpus dentatus (Aubl.) Standl., J. Wash. Acad. Sci. 15: 286 (1925). Tigarea dentata Aubl., Hist. Pl. Guiane 2: 920 (1775). Holotype: French Guiana, Aublet s.n. (BM). Illustr.: Standley y Williams, Fieldiana, Bot. 24(7): 8, t. 2 (1963). Lianas; young branches glabrous, often with grayish-white exfoliating bark. Leaves 6-24 × 3-10 cm, lanceolate to elliptic, acuminate at apex, attenuate at base, with 11-18 pairs of lateral veins, ending at margins in a sharp tooth, smooth above and beneath, leaves villous beneath along veins, coarsely dentate, non-punctate; petioles 1-2.5 cm, villous, very narrowly alate. Inflorescences axillary, fasciculate with 10-30 pedunculate flowers; peduncles 1-2.5 cm, sparsely strigose. Flowers 0.7-1.2 cm broad; sepals unequal, 2-5 mm, elliptic to obovate, pubescent externally, glabrous inside; petals c. 4 mm, obovate, white, deciduous; stamens 25-35; carpels 1, glabrous; style c. 1 mm. Fruits 0.5-0.9 cm in diameter, globose, red; seeds 1, black, with a white aril. Swamp forests, evergreen and semideciduous forests, disturbed areas. T (Ventura 19962, MO); Ch (Breedlove y Almeda 57912, MO); B (Gentle 3910, MO); G (Contreras 6096, MO); ES (Carballo et al. 1058, MO); N (Stevens 8052, MO); CR (Gómez et al. 20779, MO); P (Foster 1470, MO). 0-1000 m. (C. Mexico, Mesoamerica, Colombia, Venezuela, Guayanas, Ecuador, Perú, Bolivia, Brazil, Paraguay, Cuba, Trinidad, Tobago.) Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 9 of 15

On subspecies, spp. dentatus, occurs in Mesoamerica; six additional subspecies are found in South America.

4. Doliocarpus major J.F. Gmel., Syst. Nat. 2: 805 (1791). Holotype: Surinam, Rolander s.n. (SBT). Illustr.: not found. Lianas. Leaves 4.5-16(-20) × 2-7 cm, lanceolate to elliptic, acuminate at apex, attenuate at base, with 6-9 pairs of lateral veins, these extending to margin, smooth above and beneath, sparsely strigose beneath along veins, coarsely dentate, punctate with silica crystals; petioles 0.2-1.5 cm, strigose, very narrowly alate. Inflorescences axillary, composed of fascicles of 2-14 flowers; peduncles 0.7-2.5 cm, pubescent. Flowers 1-1.5 cm broad; sepals unequal, obovate or elliptic, outer ones 2-3 mm, inner ones 4-7 mm, pubescent externally, adpressed pilose inside; petals c. 1.5 mm, white; stamens 70-80; carpels 1, pubescent; style 0.6-0.8 cm. Fruit 1-1.3 cm in diameter, subglobose, pubescent with hairs less than 0.5 mm, red; seeds 1-2 black, with a white aril. Wet evergreen forests, seasonally semideciduous forests. 0-300 m. N (Stevens 4861, MO); CR (Liesner 2989, MO); P (Mori y Kallunki 1733, MO). (Mesoamerica, Colombia, Venezuela, Guayanas, Ecuador, Perú, Bolivia, Brazil.) One subspecies, subsp. major, occurs in Mesoamerica; the other subspecies, subsp. littoralis Kubitzki, is confined to Brazil.

5. Doliocarpus multiflorus Standl., J. Wash. Acad. Sci. 15: 285 (1925). Holotype: Panama, Standley 30285 (US). Illustr.: not found. Doliocarpus guianensis auct. non (Aubl.) Gilg. Lianas; branches glabrous, reddish-brown, with exfoliating bark. Leaves 8-20 × 3- 10 cm, elliptic to obovate, short-acuminate at apex, attenuate at base, with 6-11 pairs of lateral veins, smooth, adpressed pilose on veins beneath; petioles 1-3 cm, narrowly alate. Inflorescences 2-4 cm, axillary, fascicles of 2-4-flowered racemes or panicles, sparsely pilose; flowers 7-9 mm broad; pedicles 2-5 mm; sepals 5-6 mm, oblong to obovate, sparsely pilose externally, glabrous within; petals 3-4 mm, white; stamens 50-80; carpels 1, pilose; style 1-2 mm. Fruit up to 1.2 cm in diameter, glabrous to sparsely pilose, red; seeds 1(-2), flattened reniform, with a white aril. Wet lowland tropical forests. B (Croat Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 10 of 15

24826, MO); H (Saunders 1006, MO); N (Stevens 17737, MO); CR (Liesner et al. 15188, MO); P (Croat 13488, MO). 0-900 m. (Mexico, Mesoamerica, Colombia, Venezuela, Ecuador, Perú, Bolivia, Brazil, Cuba.)

6. Doliocarpus olivaceus Sprague et R.O. Williams ex Standl., Contr. U.S. Natl. Herb. 27: 265 (1928). Lectotype (designated by Hunter, 1965 [1966]): Panama, Maxon y Harvey 6746 (US). Illustr.: Hunter, Ann. Missouri Bot. Gard. 52: 591, t. 4 (1965). Lianas; young branches glabrous; older branches with gray exfoliating bark. Leaves 8-13 × 3.5-9 cm, elliptic, acuminate at apex, attenuate at base, with 7-11 pairs of lateral veins, mostly looped-connected with superadjacent secondaries, obscurely dentate in upper 1/4 of leaf, smooth, glabrous; petioles 1-2.5 cm, very narrowly alate, glabrous, swollen and darkened at base. Inflorescences axillary, composed of fascicles of 5-14 long-pedunculate flowers; peduncles 2-6(-12) mm. Flowers 0.9-1.3 cm broad; sepals 4-6 mm, obovate or elliptic, glabrous externally, adpressed pilose inside at base; petals c. 6 mm; stamens 40-50; carpels 1, strigose; style 4-5 mm. Fruits 1.5-2 cm, with strigose hairs 1-2 mm, red; seeds 2, with a white aril. Wet evergreen or semideciduous forests. P (Foster 846, MO). 0-300 m. (Mesoamerica, Colombia, Perú.)

5. Pinzona Mart. et Zucc. Por C.A. Todzia.

Woody lianas with simple hairs only, stems more than 30 cm circumference. Leaves alternate, petiolate, smooth, entire, stipules absent. Inflorescences paniculate; axillary flowers bisexual; sepals 3-4, subequal; petals 3, obovate; stamens numerous; carpels 2, glabrous, connate ventrally, 1-loculate; ovules 2, anatropous; styles 2, terminal; stigma peltate. Fruit baccate; seeds 1-2, aril completely surrounding seed. 1 spp. Belize to Brazil, Hispaniola, Puerto Rico, Guadeloupe, Trinidad.

1. Pinzona coriacea Mart. et Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 371 (1832). Holotype: Brazil, Martius s.n. (M). Illustr.: Aymard, Fl. Venez. Guayana 4: 682, t. 539 (1998). Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 11 of 15

Doliocarpus belizenis Lundell, D. coriaceus (Mart. et Zucc.) Gilg., D. nicaraguensis Standl. Small to large lianas; branches glabrous to strigulose. Leaves 7-16 × 6-10.5 cm, suborbicular-ovate to obovate, round or short-acuminate at apex, broadly attenuate to rounded at base, decurrent on petiole, with 7-10 pairs of lateral veins, smooth, strigulose along veins beneath; petioles 1.5-2.5 cm, alate. Inflorescences axillary, paniculate, 3-5(- 10) cm, strigulose or pilose; bracts 1-2 mm, lanceolate to ovate. Flowers c. 0.5 cm broad; sepals 3-5 mm, glabrous on both surfaces, the margin pilose outside; petals 3-5 mm, obovate, subemarginate;white to cream-colored; stamens 25-30, longer that the sepals; styles 1-2 mm. Fruits didymous, c. 5 × c. 8 mm, the locules subglobose, the locules subglobose, connate above, smooth, glabrous; seeds c. 3 mm in diameter, smooth, black with an orange aril. Lowland wet forests. B (Croat 24553, MO); H (Evans 1688, MO); N (Moreno 25553, MO); CR (Hammel y Trainer 12994, MO); P (de Nevers et al. 7570, MO). 0-400 m. (Mesoamerica, Colombia, Venezuela, Guayanas, Ecuador, Perú, Bolivia, Brazil.)

6. Tetracera L. Por C.A. Todzia y G. Aymard C.

Scandent shrubs or woody lianas, with rough (often stellate) pubescence. Leaves simple, alternate, petiolate, estipulate, coriaceous, conspicuously nerved, scabrous with the hairs simple or fasciculate. Inflorescences racemes, paniculate or thyrses, axillary or terminalmany-flowered; bracts lanceolate. Plants androdioecious. Flowers bisexual or polygamous, actinomorphic; sepals (4)5 (7), persistent, more or less equal or the outer slightly smaller; petals 3-5, caducous; stamens 50-200, persistent; carpels 1-6, pyriform, free, 1-loculed; stigmas capitate or peltate. Follicles 1-6, pyriform, dry, coriaceous, dehiscing along the ventral suture; seeds 1-4, surrounded by a lacerate aril. 45 spp. Pantropical. Literature: Kubitzki, K. Mitt. Bot. Staatssamml. München 8: 1-98 (1970).

1. Nervation craspedromous; sepals sericeous inside. Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 12 of 15

2. Leaves abruptly attenuate at the base; follicles 1. 3. T. portobellensis 2. Leaves rounded to narrowly cuneat at base; follicles 3-5. 3. Leaves glabrescent beneath; flower buds globose; follicles glabrous to sparsely strigose at the apex. 4a. T. volubilis subsp. volubilis 3. Leaves pilose beneath; flower buds obovate; follicles densely tomentose. 4b. T. volubilis subsp mollis 1. Nervation brochidodromous; sepals glabrous inside. 4. Leaves dentate. 2. T. hydrophila 4. Leaves entire or subsinuate. 5. Petioles appressed-pubescent with simple trichomes; inflorescences 2.5-3 cm; pedicels c. 3 mm; sepals 9; follicles glabrous. 1. T. macphersonii 5. Petioles stellate-pubescent; inflorescences 10-30 cm; pedicels 0.5-1 mm; sepals 5; follicles sparsely pilose at the apex. 5a. T. willdenowiana subsp. willdenowiana

1. Tetracera macphersonii G.A. Aymard, Novon 6: 325 (1996). Isotype: Panama, McPherson 9753 (MO!). Illustr.: Aymard, Novon 6: 326, t. 1 (1996). Lianas. Branches parsely stellate-pubescent, glabrescent. Leaves 3-8 × 1.5-4 cm subcoriaceous, scabrous, elliptic or obovate-elliptic, the base cuenate or obtuse, the apex rounded or acute, the margins entire, often subrevolute from the middle to the apex of the blade, the veins raised below, with 5-8 parallel nerves on each side of the mid-nerve, convergent toward margin, the surfaces lepidote-stellate above, glabrous beneath except along the midrib and secondary nerves, these sparsely appressed-pubescent; petioles 1.2- 1.7 cm, subulate, canaliculate, appressed-pubescent. Inflorescence an axillary, rarely terminal, 2-3 flowered thyrse, 2.5-3 cm, appressed-pubescent, with stellate and simple trichomes; pedicels 2-3 mm, densely stellate-pubescent. Flowers bisexual; sepals 9, suborbicular, unequal, eciliate, the outer three 4-6 mm, sparsely lepidote-stellate without, glabrous within, internal six 5-8 mm, lepidote-stellate without, glabrous within; petals not seen; stamens 80-100, 3.5-4 mm, the filaments glabrous, the anthers c. 0.5 mm. Fruit of tree follicles each 10-11 mm, coriaceous, smooth, shining and glabrous externally, opening by longitudianl slits, rostrate, the rostruym 2-3 mm; seed 1 per locule, 2-4 mm, Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 13 of 15

black, with aril longer than seed and deeply lacerate. Wet forests. P (McPherson 9753, MO). 300-400 m. (Endemic.)

2. Tetracera hydrophila Triana et Planch., Ann. Sci. Nat. Bot. sér. 4, 17: 20 (1862). Syntype: Colombia, Triana s.n. (K). Illustr.: not found. Tetracera ovalifolia auct. non DC. Lianas; young branches essentially glabrous; older branches with red exfoliating bark. Leaves 7.5-18(-22) × 4-12 cm, elliptic, round, acute to short-acuminate at apex, rounded, sometimes oblique or decurrent on petiole at base, with 9-15 pairs of lateral veins, smooth above, slightly scabrous beneath, coriaceous, margins obscurely dentate; petioles 1-2.5 cm, sparsely strigose, narrowly alate. Inflorescences terminal or axillary thyrses, 5-25 cm, stellate-pubescent. Flowers 1-1.5 cm broad; sepals c. 0.5 mm, obovate, usually glabrous, rarely sparsely pubescent inside and out; petals c. 1 cm, obovate, white; stamens 150-200; carpels 4(-5), abruptly contracted at apex, style 1-2 mm. Follicles 4-5, glabrous, light brown, 0.5-1 cm; seeds ca. 5 mm in diameter, black, glabrous. Wet evergreen forests. B (Gentry 8428, MO); H (Liesner 26405, MO); N (Araquistain 3284, MO); CR (Heithaus 210, MO); P (von Wedel 1664, MO). 0-500 m. (Mesoamerica, Colombia, Ecuador, Perú, Bolivia.)

3. Tetracera portobellensis Beurl., Kongl. Vetensk. Acad. Handl. 40: 113 (1854 [1856]). Holotype: Panama, Billberg 323 (S). Illustr.: Hunter, Ann. Missouri Bot. Gard. 52: 583, t. 1 (1966). Lianas, young branches strigose and stellate-pubescent; older branches with red exfoliating bark; young branches hirsute and stellate-pubescent. Leaves 6-20 × 4-9 cm, narrowly elliptic, elliptic to broadly elliptic, obtuse to short-acuminate at apex, abruptly attenuate at base, with 12-24 pairs of lateral nerves, scabrous and sparsely strigose above, scabrous beneath, strigose only along veins, coriaceous, margins sharply dentate; petioles 0.5-1 cm, strigose and stellate-pubescent. Inflorescences terminal or axillary, compound thyrses, 10-20 cm, sparsely villous. Flowers c. 5 mm broad; sepals suborbicular, c. 3 mm, hirsute outside, sericeous inside; petals obovate, whtie, c. 5 mm; stamens 50-80; carpels 1. essentially glabrous. Follicles 1, glabrous, dark brown, 6-9 mm; seeds c. 2.5 mm, with Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 14 of 15

a reddish-orange aril. Gallery forests, river banks, and disturbed forest edges. H (Nelson y Cruz 9331, MO); N (Neill 3721, MO); CR (Hammel 11563, MO); P (Foster 1405, MO). 0-400 m. (C. Mexico, Mesoamerica, Colombia.)

4. Tetracera volubilis L., Sp. Pl. 533 (1753). Type cons.: Mexico, Veracruz, Purpus 2206 (F). Illustr.: Standley y Williams, Fieldiana, Bot. 24(7): 11 (1963). Small shrubs, or more often, lianas, with exfoliating bark; young branches glabrous to hirsute. Leaves 7-20 × 2.5-9 cm, narrowly elliptic to narrowly obovate, round, acute to short-acuminate at apex, round to narrowly cuneat at base, with 9-15(-19) pairs of lateral veins, scabrous above and beneath, glabrous to pilose beneath, coriaceous; margins sharply dentate; petioles 1-3 cm, alate, sparsely strigose. Inflorescences axillary or terminal thyrses, 8-30 cm, pubescent. Flowers 0.5-1 cm broad; sepals 3-5 mm, orbicular, stellate-pubescent externally, seriaceous within; petals 0.5-1 cm, elliptic, white; stamens 100-200; carpels 3-5, short-rostrate; style c. 2 mm. Follicles 3-5, 0.7-1 cm, sparsely pilose at apex; seeds c. 4 mm in diameter, black, with an arange to red aril. Seasonally dry forests, low montane rain forests, premontane moist forests, disturbed forests and roadsides. T-P. 0-700 m. (Mexico, Mesoamerica, South America, Cuba, Jamaica.)

4a. Tetracera volubilis subsp. volubilis. Illustr.: Gallardo-Hernández, Fl. Veracruz 134: 25, t. 4 (2004). Leaves scabrous beneath. Inflorescence rachises sparsely stellate-pubescent; flower buds globose, 3-6 mm; sepals 3-5 mm, sparsely stellate-pubescent on outer surface. Follicles sparsely strigose at apex. T (Novelo et al. 120, MO); Ch (Méndez Ton 3808, MO); C (Lundell 1273, MO); QR (Téllez V. et al. 3619, MO); B (Davidse y Brant 32440, MO); G (Ortíz 467, MO); H (Nelson y Romero 4563, MO); ES (Morales Can 2104, MO); N (Stevens 23351, MO); CR (Liesner 4957, MO); P (Knapp 1919, MO). 0- 700 m. (Mexico [Veracruz], Mesoamerica, Colombia, Venezuela, Ecuador, Perú, Brazil, Cuba, Jamaica.)

Flora Mesoamericana, Volume 2(1), Dilleniaceae, page 15 of 15

4b. Tetracera volubilis subsp. mollis (Standl.) Kubitzki, Mitt. Bot. Staatssamml. München 8: 46 (1970). T. mollis Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 25 (1930). Holotype: Belize, Lundell 47 (F). Illustr.: not found. Leaves pilose beneath. Inflorescence rachises densely stellate-pubescent; flower buds obovate, 5-6 mm; sepals c. 7 mm, hirsute on outer surface. Follicles densely tomentose. Ch (Breedlove 26685, MO); C (Martínez et al. 27865, MO); QR (Téllez y Cabrera 3073, MO); B (Gentle 3706, MO); G (Contreras 9137, MO); P (Carrasquilla 3038, MO). 0-600 m. (Mesoamerica, Cuba.)

5. Tetracera willdenowiana Steud., Nomencl. Bot. ed. 2, 2: 670 (1841). Lectotype (designado por Kubitzki, 1970): Brazil, Hoffmannsegg s.n. (B). Illustr.: Aymard, Fl. Venez. Guayana 4: 685 (1998). Tetracera fagifoliaWilld. ex Schltdl. non Blume, T. ovalifolia auct. non DC. Lianas; young branches with exfoliating bark. Leaves 5-12 × 3.5-7.5 cm, elliptic, acute to short-acuminate, rounded to truncate to subcordate (sometimes oblique) at base, with 7-16 pairs of lateral veins, with veins joining the one above it, smooth to slightly scabrous, glabrous; margins entire; petioles 0.7-3 cm, alate only at apex, stellate- pubescent. Flowers 6-9 mm in diameter; sepals 5, orbicular, 3-4 mm, stellate-pubescent outside, glabrous within; petals c. 5 mm, obovate, white; carpels 3-4, short-rostrate; style 1-2 mm. Follicles 6-9 mm, sparsely pilose at apex; seeds c. 3 mm in diameter, black, with a red aril. Forest edges and secondary growths. G (Harmon y Dwyer 2665, MO); CR (Kernan 1071, MO); P (Antonio 2393, MO). 0-200 m. (Mesoamerica, Colombia, Venezuela, Surinam, French Guiana, Ecuador, Peru, Bolivia, Brazil.) Only subspecies. willdenowiana is known from Mesoamerica. Subspecies emarginata is restricted to Brazil and Venezuela.